Of all the evolutionary transitions that have ever taken place few have received as much attention as the origin of whales. (See here, here, here, here,and here for a few of my posts on the subject.) The story of how terrestrial hoofed mammals gave rise to the exclusively aquatic leviathans has been highlighted in headlines over and over again, but other marine mammals have not received the same amount of public attention. In the case of pinnipeds (seals, sea lions, and walruses) this may be at least partially due to the fact that their origins have been difficult to tease out.

It has long been known that seals and sea lions are carnivoran mammals closely related to weasels or bears, but just how pinnipeds first became adapted to a semi-aquatic life has been more difficult to figure out. This is largely due to the fact that their fossil trail stopped rather abruptly. The 24-22 million year old Enaliarctos, the oldest pinniped well-represented by fossils, was already a sea-lion like creature that swam in the sea. Surely there had to be even older fossils connecting it to its progenitors, but for years the trail was cold.

The skull of Puijila from the underside (a) and left side (b). From the Nature paper.

The gap between Enaliarctos and its forebears did not represent a real void in nature. Instead it pointed to a gap in our knowledge. That gap has now been partially filled with the announcement in the journal Nature of Puijila darwini, an early Miocene seal that represents a more terrestrial stage of pinniped evolution.

Bear in mind, however, that Puijila was probably not ancestral to Enaliarctos or living pinnipeds. Given that evolution is a branching process it can be extremely difficult to tell whether you are dealing with, as T.H. Huxley put it, “fathers” and “sons” (linear types) or “uncles” and “nephews” (intercalary types). Puijila appears to be one of the latter, a side branch that preserved some of the tell-tale traits than can inform us about pinniped evolutionary transitions. Indeed, the significance of Puijila is that it is representative of a stage of pinniped evolution that scientists could only hypothesize about previously. It is the fossil confirmation of the idea that seals did evolved from fully terrestrial ancestors.

Puijila was found on Devon Island, a chilly island well inside the Arctic Circle in Nunavut, Canada. About 21 million years ago there was a freshwater lake there where rabbits, rhinoceros, antelope, and other creatures sometimes came to drink.Puijila also lived in and around the lake. It was about three feet long with a long tail and broad, flat foot bones that, as in many groups of aquatic mammals, indicate that it had webbed feet. It did not have flippers, but its webbed feet certainly would have helped it get around in the water.

Overall Puijila was very similar in many respects to the extant river otters of North America, yet it differed in some important ways. It had upper arms and shoulders, for instance, that were slightly more expanded and robust than the same bones in otters. These differences would have provided more area for muscle attachment, and this could mean that Puijila was propelling itself with its webbed front and hind feet (possibly with an emphasis on the front feet).

So how does Puijila relate to other mammals? Given that it lived just prior to Enaliarctos and a radiation of marine pinnipeds it is probably more of a “persistent type” of early pinniped than an actual ancestor of creatures like Enaliarctos. Still, the phylogenetic analysis grouped it with Enaliarctos and another pinniped called Potamotherium. From what is known of these genera and the locations in which they have been found the authors of the paper suggest that pinnipeds may have originated by about 33 million years ago somewhere in the Arctic.

If the authors of the paper are correct the transition of pinnipeds to an aquatic mode of life would have started with Puijila-like mammals that lived in freshwater. Eventually, though, these amphibious mammals would have extend their range to the sea shore where they would have been further adapted to life in the water into forms more like Enaliarctos. Whether this hypothesis is correct, however, will rest on further studies of the fossil evidence.

Puijila is also important to illustrating the contingent nature of evolution. Even though it is extremely otter-like it did not swim like an otter. It primarily used its limbs to swim and, it probably did not incorporate up-and-down motions of its spinal column like transitional whales (i.e. Rodhocetus) did, either. If Puijila really does represent an important transitional stage in pinniped evolution, then, they way it swam can tell us much about why pinnipeds swim by using their limbs while other marine mammals exhibit different types of locomotion. The past is the key to the present.

I truly hope that Puijila gets the public attention it deserves. It is a wonderful, nearly-complete transitional form that answers some of our evolutionary questions while raising new ones. I hope it will inspire vertebrate paleontologists to look into pinniped origins with renewed vigor, and perhaps in a few decades we can talk about transitional pinnipeds with as much excitement as that with which we discuss transitional cetaceans.

Great write up – a bit more eloquent and less rushed than my own, I’ll admit. Thanks for linking my article too! I saw you beat me to it (I’ve been slaving away at my thesis proposal, but I’m still really excited about this cute little guy). I need to add ref’s for my article, some additional photos, etc. anyway, so I’ll return the favor.

Russel – the authors postulated webbed digits because of the flattened phalanges, which as I understand is a rough correlation among modern mammals (only from what I’ve read – I really do need to look at more carnivoran skeletons).

Webbed feet can usually be extrapolated from two osteological features: relatively flattened phalanges, and little “wide spots” at the distal ends of the terminal phalanges. This can be seen in most modern web-footed mammals, and it’s also present in Castorocauda, a Mesozoic “beaver.” These wide spots are for the attachment of the webbing, so they serve as an osteological correlate for webbed feet ‘n’ hands.

And Brian, I’d bet dollars to doughnuts that pinnipeds are foot-powered swimmers thanks to their ancestry. Their closest relatives are dogs and bears, both of which have small tails but long legs, and doggie-paddle in the water.

Excellent find. I’m thrilled to see more light being shed on the origins of yet another group of marine mammals.

It appears that many carnivoran groups had a strong affinity for a semi-aquatic or aquatic existence. Besides the lineage that gave rise to the pinnipeds, there’s also the bears – polar bear and the Miocene Kolponomos. Among the mustelids, there are otters and mink (2 lineages? If American mink and sea mink are a different genus from European mink). The civets have the otter genet and the otter civet. Raccoons seem strongly tied to water as well. There is a marsh mongoose. And the fishing cat. It seems that various carnivore lineages have evolved to live along the water’s edge time and time again.

This is my favorite intermediate find since the land-walking sirenian Pezosiren. Your point about the “branching bush” and Puijilla not being ancestral to modern pinnipeds is well taken. John Wilkins on his blog attacked the whole concept of “missing links”, and the simplistic picture of evolution such concepts paint. If an Archaeopteryx isn’t ancestral to modern birds, or Puijila ancestral to modern seals or sea lions, it’s no less of an intermediary for that.

This is my favorite intermediate find since the land-walking sirenian Pezosiren. Your point about the “branching bush” and Puijilla not being ancestral to modern pinnipeds is well taken. John Wilkins on his blog attacked the whole concept of “missing links”, and the simplistic picture of evolution such concepts paint. If an Archaeopteryx isn’t anc

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